[pointprocesses] improved chapter

This commit is contained in:
Jan Benda 2021-01-18 13:13:35 +01:00
parent 0f0dfafd56
commit 20332e2013
10 changed files with 272 additions and 194 deletions

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@ -1,16 +1,11 @@
function [counts, bins] = counthist(spikes, w) function counthist(spikes, w)
% Compute and plot histogram of spike counts. % Plot histogram of spike counts.
% %
% [counts, bins] = counthist(spikes, w) % counthist(spikes, w)
% %
% Arguments: % Arguments:
% spikes: a cell array of vectors of spike times in seconds % spikes: a cell array of vectors of spike times in seconds
% w: observation window duration in seconds for computing the counts % w: duration of window in seconds for computing the counts
%
% Returns:
% counts: the histogram of counts normalized to probabilities
% bins: the bin centers for the histogram
% collect spike counts: % collect spike counts:
tmax = spikes{1}(end); tmax = spikes{1}(end);
n = []; n = [];
@ -21,12 +16,12 @@ function [counts, bins] = counthist(spikes, w)
% for tk = 0:w:tmax-w % for tk = 0:w:tmax-w
% nn = sum((times >= tk) & (times < tk+w)); % nn = sum((times >= tk) & (times < tk+w));
% %nn = length(find((times >= tk) & (times < tk+w))); % %nn = length(find((times >= tk) & (times < tk+w)));
% n = [n nn]; % n = [n, nn];
% end % end
% alternative 2: use the hist() function to do that! % alternative 2: use the hist() function to do that!
tbins = 0.5*w:w:tmax-0.5*w; tbins = 0.5*w:w:tmax-0.5*w;
nn = hist(times, tbins); nn = hist(times, tbins);
n = [n nn]; n = [n, nn];
end end
% histogram of spike counts: % histogram of spike counts:
@ -36,9 +31,7 @@ function [counts, bins] = counthist(spikes, w)
counts = counts / sum(counts); counts = counts / sum(counts);
% plot: % plot:
if nargout == 0
bar(bins, counts); bar(bins, counts);
xlabel('counts k'); xlabel('counts k');
ylabel('P(k)'); ylabel('P(k)');
end end
end

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@ -8,7 +8,7 @@ function [pdf, centers] = isihist(isis, binwidth)
% binwidth: optional width in seconds to be used for the isi bins % binwidth: optional width in seconds to be used for the isi bins
% %
% Returns: % Returns:
% pdf: vector with probability density of interspike intervals in Hz % pdf: vector with pdf of interspike intervals in Hertz
% centers: vector with centers of interspikeintervalls in seconds % centers: vector with centers of interspikeintervalls in seconds
if nargin < 2 if nargin < 2

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@ -5,11 +5,9 @@ function isivec = isis(spikes)
% %
% Arguments: % Arguments:
% spikes: a cell array of vectors of spike times in seconds % spikes: a cell array of vectors of spike times in seconds
% isivec: a column vector with all the interspike intervalls
% %
% Returns: % Returns:
% isivec: a column vector with all the interspike intervalls % isivec: a column vector with all the interspike intervalls
isivec = []; isivec = [];
for k = 1:length(spikes) for k = 1:length(spikes)
difftimes = diff(spikes{k}); difftimes = diff(spikes{k});

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@ -1,15 +1,15 @@
function isicorr = isiserialcorr(isivec, maxlag) function [isicorr, lags] = isiserialcorr(isivec, maxlag)
% serial correlation of interspike intervals % serial correlation of interspike intervals
% %
% isicorr = isiserialcorr(isivec, maxlag) % isicorr = isiserialcorr(isivec, maxlag)
% %
% Arguments: % Arguments:
% isivec: vector of interspike intervals in seconds % isivec: vector of interspike intervals in seconds
% maxlag: the maximum lag in seconds % maxlag: the maximum lag
% %
% Returns: % Returns:
% isicorr: vector with the serial correlations for lag 0 to maxlag % isicorr: vector with the serial correlations for lag 0 to maxlag
% lags: vector with the lags corresponding to isicorr
lags = 0:maxlag; lags = 0:maxlag;
isicorr = zeros(size(lags)); isicorr = zeros(size(lags));
for k = 1:length(lags) for k = 1:length(lags)
@ -21,14 +21,4 @@ function isicorr = isiserialcorr(isivec, maxlag)
isicorr(k) = corr(isivec(1:end-lag), isivec(lag+1:end)); isicorr(k) = corr(isivec(1:end-lag), isivec(lag+1:end));
end end
end end
if nargout == 0
% plot:
plot(lags, isicorr, '-b');
hold on;
scatter(lags, isicorr, 50.0, 'b', 'filled');
hold off;
xlabel('Lag k')
ylabel('\rho_k')
end
end end

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@ -0,0 +1,16 @@
function isicorr = plotisiserialcorr(isivec, maxlag)
% plot serial correlation of interspike intervals
%
% plotisiserialcorr(isivec, maxlag)
%
% Arguments:
% isivec: vector of interspike intervals in seconds
% maxlag: the maximum lag
[isicorr, lags] = isiserialcorr(isivec, maxlag);
plot(lags, isicorr, '-b');
hold on;
scatter(lags, isicorr, 20.0, 'b', 'filled');
hold off;
xlabel('Lag k')
ylabel('\rho_k')
end

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@ -2,21 +2,35 @@ function rasterplot(spikes, tmax)
% Display a spike raster of the spike times given in spikes. % Display a spike raster of the spike times given in spikes.
% %
% rasterplot(spikes, tmax) % rasterplot(spikes, tmax)
%
% Arguments:
% spikes: a cell array of vectors of spike times in seconds % spikes: a cell array of vectors of spike times in seconds
% tmax: plot spike raster up to tmax seconds % tmax: plot spike raster up to tmax seconds
in_msecs = tmax < 1.5
spiketimes = [];
trials = [];
ntrials = length(spikes); ntrials = length(spikes);
for k = 1:ntrials for k = 1:ntrials
times = spikes{k}; times = spikes{k};
times = times(times<tmax); times = times(times<tmax);
if tmax < 1.5 if in_msecs
times = 1000.0*times; % conversion to ms times = 1000.0*times; % conversion to ms
end end
for i = 1:length( times ) % (x,y) pairs for start and stop of stroke
line([times(i) times(i)],[k-0.4 k+0.4], 'Color', 'k'); % plus nan separating strokes:
end spiketimes = [spiketimes, ...
[times(:)'; times(:)'; times(:)'*nan]];
trials = [trials, ...
[ones(1, length(times)) * (k-0.4); ...
ones(1, length(times)) * (k+0.4); ...
ones(1, length(times)) * nan]];
end end
if tmax < 1.5 % convert matrices into simple vectors of (x,y) pairs:
spiketimes = spiketimes(:);
trials = trials(:);
% plotting this is lightning fast:
plot(spiketimes, trials, 'k')
if in_msecs
xlabel('Time [ms]'); xlabel('Time [ms]');
xlim([0.0 1000.0*tmax]); xlim([0.0 1000.0*tmax]);
else else

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@ -92,21 +92,37 @@ def plot_count_fano(ax1, ax2, spikes):
ax2.set_yticks(np.arange(0.0, 1.2, 0.5)) ax2.set_yticks(np.arange(0.0, 1.2, 0.5))
def plot_fano(ax, spikes):
wins = np.logspace(-2, 0.0, 200)
mean_counts = np.zeros(len(wins))
var_counts = np.zeros(len(wins))
for k, win in enumerate(wins):
counts = []
for times in spikes:
c, _ = np.histogram(times, np.arange(0.0, duration, win))
counts.extend(c)
mean_counts[k] = np.mean(counts)
var_counts[k] = np.var(counts)
ax.plot(1000.0*wins, var_counts/mean_counts, **lsB)
ax.set_xlabel('Window', 'ms')
ax.set_ylim(0.0, 1.2)
ax.set_xscale('log')
ax.set_xticks([10, 100, 1000])
ax.set_xticklabels(['10', '100', '1000'])
ax.xaxis.set_minor_locator(mpt.NullLocator())
ax.set_yticks(np.arange(0.0, 1.2, 0.5))
if __name__ == "__main__": if __name__ == "__main__":
homspikes = hompoisson(rate, trials, duration) homspikes = hompoisson(rate, trials, duration)
inhspikes = oupifspikes(rate, trials, duration, dt, 0.3, drate, tau) inhspikes = oupifspikes(rate, trials, duration, dt, 0.3, drate, tau)
fig, axs = plt.subplots(2, 2) fig, (ax1, ax2) = plt.subplots(1, 2)
fig.subplots_adjust(**adjust_fs(fig, top=0.5, right=2.0)) fig.subplots_adjust(**adjust_fs(fig, top=0.5, right=2.0))
plot_count_fano(axs[0,0], axs[0,1], homspikes) plot_fano(ax1, homspikes)
axs[0,0].text(0.1, 0.95, 'Poisson', transform=axs[0,0].transAxes) ax1.set_ylabel('Fano factor')
axs[0,0].set_xlabel('') ax1.text(0.1, 0.95, 'Poisson', transform=ax1.transAxes)
axs[0,1].set_xlabel('') plot_fano(ax2, inhspikes)
axs[0,0].xaxis.set_major_formatter(mpt.NullFormatter()) ax2.axhline(1.0, **lsGrid)
axs[0,1].xaxis.set_major_formatter(mpt.NullFormatter()) ax2.text(0.1, 0.95, 'OU noise', transform=ax2.transAxes)
plot_count_fano(axs[1,0], axs[1,1], inhspikes)
axs[1,1].axhline(1.0, **lsGrid)
axs[1,0].text(0.1, 0.95, 'OU noise', transform=axs[1,0].transAxes)
fig.text(0.01, 0.58, 'Count variance', va='center', rotation='vertical')
fig.text(0.51, 0.58, 'Fano factor', va='center', rotation='vertical')
plt.savefig('fanoexamples.pdf') plt.savefig('fanoexamples.pdf')
plt.close() plt.close()

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@ -13,9 +13,9 @@ information. Analyzing the statistics of spike times and their
relation to sensory stimuli or motor actions is central to relation to sensory stimuli or motor actions is central to
neuroscientific research. With multi-electrode arrays it is nowadays neuroscientific research. With multi-electrode arrays it is nowadays
possible to record from hundreds or even thousands of neurons possible to record from hundreds or even thousands of neurons
simultaneously. The open challenge is how to analyze such data sets in simultaneously. The open challenge is how to analyze such huge data
order to understand how neural systems work. Let's start with the sets in smart ways in order to gain insights into the way neural
basics in this chapter. systems work. Let's start with the basics in this chapter.
The result of the pre-processing of electrophysiological recordings The result of the pre-processing of electrophysiological recordings
are series of spike times, which are termed \enterm[spike train]{spike are series of spike times, which are termed \enterm[spike train]{spike
@ -27,15 +27,16 @@ process]{Punktprozess}{point processes}.
\begin{figure}[bt] \begin{figure}[bt]
\includegraphics[width=1\textwidth]{rasterexamples} \includegraphics[width=1\textwidth]{rasterexamples}
\titlecaption{\label{rasterexamplesfig}Raster plot.}{Raster plots of \titlecaption{\label{rasterexamplesfig}Raster plots of spike
ten trials of data illustrating the times of action trains.}{Raster plots of ten trials of data illustrating the times
potentials. Each vertical stroke illustrates the time at which an of action potentials. Each vertical stroke illustrates the time at
action potential was observed. Each row displays the events of one which an action potential was observed. Each row displays the
trial. Shown is a stationary point process (homogeneous point events of one trial. Shown is a stationary point process
process with a rate $\lambda=20$\;Hz, left) and an non-stationary (homogeneous point process with a rate $\lambda=20$\;Hz, left) and
point process with a rate that varies in time (noisy perfect an non-stationary point process with a rate that varies in time
integrate-and-fire neuron driven by Ornstein-Uhlenbeck noise with (noisy perfect integrate-and-fire neuron driven by
a time-constant $\tau=100$\,ms, right).} Ornstein-Uhlenbeck noise with a time-constant $\tau=100$\,ms,
right).}
\end{figure} \end{figure}
@ -59,9 +60,10 @@ process]{Punktprozess}{point processes}.
\begin{figure}[tb] \begin{figure}[tb]
\includegraphics{pointprocesssketch} \includegraphics{pointprocesssketch}
\titlecaption{\label{pointprocesssketchfig} Statistics of point \titlecaption{\label{pointprocesssketchfig} Statistics of point
processes.}{A point process is a sequence of instances in time processes.}{A temporal point process is a sequence of events in
$t_i$ that can be also characterized by inter-event intervals time, $t_i$, that can be also characterized by the corresponding
$T_i=t_{i+1}-t_i$ and event counts $n_i$.} inter-event intervals $T_i=t_{i+1}-t_i$ and event counts $n_i$,
i.e. the number of events that occurred so far.}
\end{figure} \end{figure}
\noindent \noindent
@ -77,75 +79,108 @@ plot in which each vertical line indicates the time of an event. The
event from two different point processes are shown in event from two different point processes are shown in
\figref{rasterexamplesfig}. In addition to the event times, point \figref{rasterexamplesfig}. In addition to the event times, point
processes can be described using the intervals $T_i=t_{i+1}-t_i$ processes can be described using the intervals $T_i=t_{i+1}-t_i$
between successive events or the number of observed events within a between successive events or the number of observed events $n_i$
certain time window $n_i$ (\figref{pointprocesssketchfig}). within a certain time window (\figref{pointprocesssketchfig}).
\begin{exercise}{rasterplot.m}{} In \enterm[point process!stationary]{stationary} point processes the
Implement a function \varcode{rasterplot()} that displays the times of statistics does not change over time. In particular, the rate of the
action potentials within the first \varcode{tmax} seconds in a raster process, the number of events per time, is constant. The homogeneous
plot. The spike times (in seconds) recorded in the individual trials point process shown in \figref{rasterexamplesfig} on the left is an
are stored as vectors of times within a cell array. example of a stationary point process. Although locally within each
\end{exercise} trial there are regions with many events per time and others with long
intervals between events, the average number of events within a small
time window over trials does not change in time. In the first sections
of this chapter we introduce various statistics for characterizing
stationary point processes.
On the other hand, the example shown in \figref{rasterexamplesfig} on
the right is a non-stationary point process. The rate of the events in
all trials first decreases and then increases again. This common
change in rate may have been caused by some sensory input to the
neuron. How to estimate temporal changes in event rates (firing rates)
is covered in section~\ref{nonstationarysec}.
%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%% %%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%
\section{Homogeneous Poisson process} \section{Homogeneous Poisson process}
The Gaussian distribution is, because of the central limit theorem, Before we are able to start analyzing point processes, we need some
the standard distribution for continuous measures. The equivalent in data. As before, because we are working with computers, we can easily
the realm of point processes is the simulate them. To get us started we here briefly introduce the
\entermde[distribution!Poisson]{Verteilung!Poisson-}{Poisson distribution}. homogeneous Poisson process. The Poisson process is to point processes
what the Gaussian distribution is to the statistics of real-valued
data. It is a standard against which everything is compared.
In a \entermde[Poisson process!homogeneous]{Poissonprozess!homogener}{homogeneous Poisson In a \entermde[Poisson
process} the events occur at a fixed rate $\lambda=\text{const}$ and process!homogeneous]{Poissonprozess!homogener}{homogeneous Poisson
are independent of both the time $t$ and occurrence of previous events process} events at every given time, that is within every small time
(\figref{hompoissonfig}). The probability of observing an event within window of width $\Delta t$ occur with the same constant
a small time window of width $\Delta t$ is given by probability. This probability is independent of absolute time and
\begin{equation} independent of any events occurring before. To observe an event right
\label{hompoissonprob} after an event is as likely as to observe an event at some specific
P = \lambda \cdot \Delta t \; . time later on.
\end{equation}
In an \entermde[Poisson process!inhomogeneous]{Poissonprozess!inhomogener}{inhomogeneous Poisson The simplest way to simulate events of a homogeneous Poisson process
process}, however, the rate $\lambda$ depends on time: $\lambda = is based on the exponential distribution \eqref{hompoissonexponential}
\lambda(t)$. of event intervals. We randomly draw intervals from this distribution
and then sum them up to convert them to event times. The only
\begin{exercise}{poissonspikes.m}{} parameter of a homogeneous Poisson process is its rate. It defines how
Implement a function \varcode{poissonspikes()} that uses a homogeneous many events per time are expected.
Poisson process to generate events at a given rate for a certain
duration and a number of trials. The rate should be given in Hertz Here is a function that generates several trials of a homogeneous
and the duration of the trials is given in seconds. The function Poisson process:
should return the event times in a cell-array. Each entry in this
array represents the events observed in one trial. Apply \lstinputlisting[caption={hompoissonspikes.m}]{hompoissonspikes.m}
\eqnref{hompoissonprob} to generate the event times.
\end{exercise}
%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%
\section{Raster plot}
Let's generate some events with this function and display them in a
raster plot as in \figref{rasterexamplesfig}. For the raster plot we
need to draw for each event a line at the corresponding time of the
event and at the height of the corresponding trial. You can go with a
one for-loop through the trials and then with another for-loop through
the event times and every time plot a two-point line. This, however,
is very slow. The fastest way is to concatenate the coordinates of all
strokes into large vectors, separate the events by \varcode{nan}
entries, and pass this to a single call of \varcode{plot()}:
\pageinputlisting[caption={rasterplot.m}]{rasterplot.m}
Adapt this function to your needs and use it where ever possible to
illustrate your spike train data to your readers. They appreciate
seeing your raw data and being able to judge the data for themselves
before you go on analyzing firing rates, interspike-interval
correlations, etc.
\begin{exercise}{hompoissonspikes.m}{}
Implement a function \varcode{hompoissonspikes()} that uses a
homogeneous Poisson process to generate spike events at a given rate
for a certain duration and a number of trials. The rate should be
given in Hertz and the duration of the trials is given in
seconds. The function should return the event times in a
cell-array. Each entry in this array represents the events observed
in one trial. Apply \eqnref{poissonintervals} to generate the event
times.
\end{exercise}
%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%% %%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%
\section{Interval statistics} \section{Interval statistics}
The intervals $T_i=t_{i+1}-t_i$ between successive events are real Let's start with the interval statistics of stationary point
positive numbers. In the context of action potentials they are processes. The intervals $T_i=t_{i+1}-t_i$ between successive events
referred to as \entermde[interspike are real positive numbers. In the context of action potentials they
are referred to as \entermde[interspike
interval]{Interspikeintervall}{interspike intervals}, in short interval]{Interspikeintervall}{interspike intervals}, in short
\entermde[ISI|see{interspike \entermde[ISI|see{interspike
interval}]{ISI|see{Interspikeintervall}}{ISI}s. The statistics of interval}]{ISI|see{Interspikeintervall}}{ISI}s. For analyzing event
interspike intervals are described using common measures for intervals we can use all the usual statistics that we know from
describing the statistics of real-valued variables: describing univariate data sets of real numbers:
\begin{figure}[t] \begin{figure}[t]
\includegraphics[width=0.96\textwidth]{isihexamples}\vspace{-2ex} \includegraphics[width=0.96\textwidth]{isihexamples}
\titlecaption{\label{isihexamplesfig}Interspike-interval \titlecaption{\label{isihexamplesfig}Interspike-interval
histograms}{of the spike trains shown in \figref{rasterexamplesfig}.} histograms}{of the spike trains shown in
\figref{rasterexamplesfig}. The intervals of the homogeneous
Poisson process (left) are exponentially distributed according to
\eqnref{hompoissonexponential} (red). Typically for many sensory
neurons that fire more regularly is an ISI histogram like the one
shown on the right. There is a preferred interval where the
distribution peaks. The distribution falls off quickly towards
smaller intervals, and very small intervals are absent, probably
because of refractoriness of the spike generator. The tail of the
distribution again approaches an exponential distribution as for
the Poisson process.}
\end{figure} \end{figure}
\begin{exercise}{isis.m}{} \begin{exercise}{isis.m}{}
@ -168,19 +203,25 @@ describing the statistics of real-valued variables:
p_{exp}(T) = \lambda e^{-\lambda T} p_{exp}(T) = \lambda e^{-\lambda T}
\end{equation} \end{equation}
of a homogeneous Poisson spike train with rate $\lambda$. of a homogeneous Poisson spike train with rate $\lambda$.
\item Mean interval: $\mu_{ISI} = \langle T \rangle = \item Mean interval
\frac{1}{n}\sum\limits_{i=1}^n T_i$. The average time it takes from \begin{equation}
one event to the next. The inverse of the mean interval is identical \label{meanisi}
with the mean rate $\lambda$ (number of events per time, see below) \mu_{ISI} = \langle T \rangle = \frac{1}{n}\sum_{i=1}^n T_i
of the process. \end{equation}
\item Standard deviation of intervals: $\sigma_{ISI} = \sqrt{\langle The average time it takes from one event to the next. For stationary
(T - \langle T \rangle)^2 \rangle}$. Periodically spiking neurons point processes the inverse of the mean interval is identical with
have little variability in their intervals, whereas many cortical the mean rate $\lambda$ (number of events per time, see below) of
neurons cover a wide range with their intervals. The standard the process.
deviation of homogeneous Poisson spike trains, $\sigma_{ISI} = \item Standard deviation of intervals
\frac{1}{\lambda}$, also equals the inverse rate. Whether the \begin{equation}
standard deviation of intervals is low or high, however, is better \label{stdisi}
quantified by the \sigma_{ISI} = \sqrt{\langle (T - \langle T \rangle)^2 \rangle}
\end{equation}
Periodically spiking neurons have little variability in their
intervals, whereas many cortical neurons cover a wide range with
their intervals. The standard deviation of homogeneous Poisson spike
trains also equals the inverse rate. Whether the standard deviation
of intervals is low or high, however, is better quantified by the
\item \entermde[coefficient of \item \entermde[coefficient of
variation]{Variationskoeffizient}{Coefficient of variation}, the variation]{Variationskoeffizient}{Coefficient of variation}, the
standard deviation of the ISIs relative to their mean: standard deviation of the ISIs relative to their mean:
@ -188,28 +229,29 @@ describing the statistics of real-valued variables:
\label{cvisi} \label{cvisi}
CV_{ISI} = \frac{\sigma_{ISI}}{\mu_{ISI}} CV_{ISI} = \frac{\sigma_{ISI}}{\mu_{ISI}}
\end{equation} \end{equation}
Homogeneous Poisson spike trains have an CV of exactly one. The Homogeneous Poisson spike trains have an $CV$ of exactly one. The
lower the CV the more regularly firing a neuron is firing. CVs lower the $CV$ the more regularly a neuron is firing. $CV$s larger than
larger than one are also possible in spike trains with small one are also possible in spike trains with small intervals separated
intervals separated by really long ones. by really long ones.
%\item \entermde[diffusion coefficient]{Diffusionskoeffizient}{Diffusion coefficient}: $D_{ISI} = %\item \entermde[diffusion coefficient]{Diffusionskoeffizient}{Diffusion coefficient}: $D_{ISI} =
% \frac{\sigma_{ISI}^2}{2\mu_{ISI}^3}$. % \frac{\sigma_{ISI}^2}{2\mu_{ISI}^3}$.
\end{itemize} \end{itemize}
\begin{exercise}{isihist.m}{} \begin{exercise}{isihist.m}{}
Implement a function \varcode{isiHist()} that calculates the normalized Implement a function \varcode{isihist()} that calculates the
interspike interval histogram. The function should take two input normalized interspike interval histogram. The function should take
arguments; (i) a vector of interspike intervals and (ii) the width two input arguments; (i) a vector of interspike intervals and (ii)
of the bins used for the histogram. It further returns the the width of the bins used for the histogram. It returns the
probability density as well as the centers of the bins. probability density as well as the centers of the bins.
\end{exercise} \end{exercise}
\begin{exercise}{plotisihist.m}{} \begin{exercise}{plotisihist.m}{}
Implement a function that takes the return values of Implement a function that takes the returned values of
\varcode{isiHist()} as input arguments and then plots the data. The \varcode{isihist()} as input arguments and then plots the data. The
plot should show the histogram with the x-axis scaled to plot should show the histogram with the x-axis scaled to
milliseconds and should be annotated with the average ISI, the milliseconds and should be annotated with the average ISI, the
standard deviation and the coefficient of variation. standard deviation, and the coefficient of variation of the ISIs
(\figref{isihexamplesfig}).
\end{exercise} \end{exercise}
\subsection{Interval correlations} \subsection{Interval correlations}
@ -227,19 +269,20 @@ correlation coefficients. We form $(x,y)$ data pairs by taking the
series of intervals $T_i$ as $x$-data values and pairing them with the series of intervals $T_i$ as $x$-data values and pairing them with the
$k$-th next intervals $T_{i+k}$ as $y$-data values. The parameter $k$ $k$-th next intervals $T_{i+k}$ as $y$-data values. The parameter $k$
is called \enterm{lag} (\determ{Verz\"ogerung}). For lag one we pair is called \enterm{lag} (\determ{Verz\"ogerung}). For lag one we pair
each interval with the next one. A \entermde[return map]{return each interval with the next one. A \entermde{return map}{return map}
map}{Return map} illustrates dependencies between successive illustrates dependencies between successive intervals by simply
intervals by simply plotting the intervals $T_{i+k}$ against the plotting the intervals $T_{i+k}$ against the intervals $T_i$ in a
intervals $T_i$ in a scatter plot (\figref{returnmapfig}). For Poisson scatter plot (\figref{returnmapfig}). For Poisson spike trains there
spike trains there is no structure beyond the one expected from the is no structure beyond the one expected from the exponential
exponential interspike interval distribution, hinting at neighboring interspike interval distribution, hinting at neighboring interspike
interspike intervals being independent of each other. For the spike intervals being independent of each other. For the spike train based
train based on an Ornstein-Uhlenbeck process the return map is more on an Ornstein-Uhlenbeck process the return map is more clustered
clustered along the diagonal, hinting at a positive correlation along the diagonal, hinting at a positive correlation between
between succeeding intervals. That is, short intervals are more likely succeeding intervals. That is, short intervals are more likely to be
to be followed by short ones and long intervals more likely by long followed by short ones and long intervals more likely by long
ones. This temporal structure was already clearly visible in the spike ones. This temporal structure was already clearly visible in each
raster shown in \figref{rasterexamplesfig}. trial of the spike raster shown in \figref{rasterexamplesfig} on the
right.
\begin{figure}[tp] \begin{figure}[tp]
\includegraphics[width=1\textwidth]{serialcorrexamples} \includegraphics[width=1\textwidth]{serialcorrexamples}
@ -259,17 +302,17 @@ raster shown in \figref{rasterexamplesfig}.
Such dependencies can be further quantified by Such dependencies can be further quantified by
\entermde[correlation!serial]{Korrelation!serielle}{serial \entermde[correlation!serial]{Korrelation!serielle}{serial
correlations}. These are the correlation coefficients between the correlations}. These quantify the correlations between successive
intervals $T_{i+k}$ and $T_i$ in dependence on lag $k$: intervals by Pearson's correlation coefficients between the intervals
$T_{i+k}$ and $T_i$ in dependence on lag $k$:
\begin{equation} \begin{equation}
\label{serialcorrelation} \label{serialcorrelation}
\rho_k = \frac{\langle (T_{i+k} - \langle T \rangle)(T_i - \langle T \rangle) \rangle}{\langle (T_i - \langle T \rangle)^2\rangle} = \frac{{\rm cov}(T_{i+k}, T_i)}{{\rm var}(T_i)} \rho_k = \frac{\langle (T_{i+k} - \langle T \rangle)(T_i - \langle T \rangle) \rangle}{\langle (T_i - \langle T \rangle)^2\rangle} = \frac{{\rm cov}(T_{i+k}, T_i)}{{\rm var}(T_i)}
= {\rm corr}(T_{i+k}, T_i) = {\rm corr}(T_{i+k}, T_i)
\end{equation} \end{equation}
The serial correlations $\rho_k$ are usually plotted against the lag The serial correlations $\rho_k$ are usually plotted against the lag
$k$ for a range small range of lags $k$ for a small range of lags (\figref{returnmapfig}). $\rho_0=1$ is
(\figref{returnmapfig}). $\rho_0=1$ is the correlation of each the correlation of each interval with itself and always equals one.
interval with itself and always equals one.
If the serial correlations all equal zero, $\rho_k =0$ for $k>0$, then If the serial correlations all equal zero, $\rho_k =0$ for $k>0$, then
the length of an interval is independent of all the previous the length of an interval is independent of all the previous
@ -282,15 +325,21 @@ thus spike trains may approximate renewal processes.
However, other variables like the intracellular calcium concentration However, other variables like the intracellular calcium concentration
or the states of slowly switching ion channels may carry information or the states of slowly switching ion channels may carry information
from one interspike interval to the next and thus introducing from one interspike interval to the next and thus introduce
correlations. Such non-renewal dynamics can then be described by the correlations between intervals. Such non-renewal dynamics is then
non-zero serial correlations (\figref{returnmapfig}). characterized by the non-zero serial correlations
(\figref{returnmapfig}).
\begin{exercise}{isiserialcorr.m}{} \begin{exercise}{isiserialcorr.m}{}
Implement a function \varcode{isiserialcorr()} that takes a vector of Implement a function \varcode{isiserialcorr()} that takes a vector
interspike intervals as input argument and calculates the serial of interspike intervals as input argument and calculates the serial
correlation. The function should further plot the serial correlations up to some maximum lag.
correlation. \end{exercise}
\begin{exercise}{plotisiserialcorr.m}{}
Implement a function \varcode{plotisiserialcorr()} that takes a
vector of interspike intervals as input argument and generates a
plot of the serial correlations.
\end{exercise} \end{exercise}
@ -326,10 +375,11 @@ split into many segments $i$, each of duration $W$, and the number of
events $n_i$ in each of the segments can be counted. The integer event events $n_i$ in each of the segments can be counted. The integer event
counts can be quantified in the usual ways: counts can be quantified in the usual ways:
\begin{itemize} \begin{itemize}
\item Histogram of the counts $n_i$. For homogeneous Poisson spike \item Histogram of the counts $n_i$ appropriately normalized to
trains with rate $\lambda$ the resulting probability distributions probability distributions. For homogeneous Poisson spike trains with
follow a Poisson distribution (\figref{countstatsfig}), where the rate $\lambda$ the resulting probability distributions follow a
probability of counting $k$ events within a time window $W$ is given by Poisson distribution (\figref{countstatsfig}), where the probability
of counting $k$ events within a time window $W$ is given by
\begin{equation} \begin{equation}
\label{poissondist} \label{poissondist}
P(k) = \frac{(\lambda W)^k e^{\lambda W}}{k!} P(k) = \frac{(\lambda W)^k e^{\lambda W}}{k!}
@ -338,7 +388,7 @@ counts can be quantified in the usual ways:
\item Variance of counts: \item Variance of counts:
$\sigma_n^2 = \langle (n - \langle n \rangle)^2 \rangle$. $\sigma_n^2 = \langle (n - \langle n \rangle)^2 \rangle$.
\end{itemize} \end{itemize}
Because spike counts are unitless and positive numbers, the Because spike counts are unitless and positive numbers the
\begin{itemize} \begin{itemize}
\item \entermde{Fano Faktor}{Fano factor} (variance of counts divided \item \entermde{Fano Faktor}{Fano factor} (variance of counts divided
by average count) by average count)
@ -351,7 +401,6 @@ Because spike counts are unitless and positive numbers, the
homogeneous Poisson processes the Fano factor equals one, homogeneous Poisson processes the Fano factor equals one,
independently of the time window $W$. independently of the time window $W$.
\end{itemize} \end{itemize}
is an additional measure quantifying event counts.
Note that all of these statistics depend in general on the chosen Note that all of these statistics depend in general on the chosen
window length $W$. The average spike count, for example, grows window length $W$. The average spike count, for example, grows
@ -372,33 +421,33 @@ information encoded in the mean spike count is transmitted.
\begin{figure}[t] \begin{figure}[t]
\includegraphics{fanoexamples} \includegraphics{fanoexamples}
\titlecaption{\label{fanofig} \titlecaption{\label{fanofig} Fano factor.}{Counting events in time
Count variance and Fano factor.}{Variance of event counts as a windows of given duration and then dividing the variance of the
function of mean counts obtained by varying the duration of the counts by their mean results in the Fano factor. Here, the Fano
count window (left). Dividing the count variance by the respective factor is plotted as a function of the duration of the window used
mean results in the Fano factor that can be plotted as a function to count events. For Poisson spike trains the variance always
of the count window (right). For Poisson spike trains the variance equals the mean counts and consequently the Fano factor equals one
always equals the mean counts and consequently the Fano factor irrespective of the count window (left). A spike train with
equals one irrespective of the count window (top). A spike train positive correlations between interspike intervals (caused by an
with positive correlations between interspike intervals (caused by Ornstein-Uhlenbeck process) has a minimum in the Fano factor, that
an Ornstein-Uhlenbeck process) has a minimum in the Fano factor, is an analysis window for which the relative count variance is
that is an analysis window for which the relative count variance minimal somewhere close to the correlation time scale of the
is minimal somewhere close to the correlation time scale of the interspike intervals (right).}
interspike intervals (bottom).}
\end{figure} \end{figure}
\begin{exercise}{counthist.m}{} \begin{exercise}{counthist.m}{}
Implement a function \varcode{counthist()} that calculates and plots Implement a function \varcode{counthist()} that calculates and plots
the distribution of spike counts observed in a certain time the distribution of spike counts observed in a certain time
window. The function should take two input arguments: (i) a window. The function should take two input arguments: a cell-array
cell-array of vectors containing the spike times in seconds observed of vectors containing the spike times in seconds observed in a
in a number of trials, and (ii) the duration of the time window that number of trials, and the duration of the time window that is used
is used to evaluate the counts. to evaluate the counts.
\end{exercise} \end{exercise}
%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%% %%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%
\section{Time-dependent firing rate} \section{Time-dependent firing rate}
\label{nonstationarysec}
So far we have discussed stationary spike trains. The statistical properties So far we have discussed stationary spike trains. The statistical properties
of these did not change within the observation time (stationary point of these did not change within the observation time (stationary point
@ -566,6 +615,7 @@ relevate time-scale.
\end{exercise} \end{exercise}
\section{Spike-triggered Average} \section{Spike-triggered Average}
\label{stasec}
The graphical representation of the neuronal activity alone is not The graphical representation of the neuronal activity alone is not
sufficient tot investigate the relation between the neuronal response sufficient tot investigate the relation between the neuronal response

View File

@ -85,7 +85,7 @@ def plot_inhomogeneous_spikes(ax):
if __name__ == "__main__": if __name__ == "__main__":
fig, (ax1, ax2) = plt.subplots(1, 2, figsize=cm_size(figure_width, 0.5*figure_width)) fig, (ax1, ax2) = plt.subplots(1, 2)
fig.subplots_adjust(**adjust_fs(fig, left=4.0, right=1.0, top=1.2)) fig.subplots_adjust(**adjust_fs(fig, left=4.0, right=1.0, top=1.2))
plot_homogeneous_spikes(ax1) plot_homogeneous_spikes(ax1)
plot_inhomogeneous_spikes(ax2) plot_inhomogeneous_spikes(ax2)

View File

@ -116,9 +116,10 @@
\includechapter{pointprocesses} \includechapter{pointprocesses}
% add a chapter on simulating point-processes/spike trains % add a chapter on simulating point-processes/spike trains
% (Poisson spike trains, integrate-and-fire models) % (Poisson spike trains, LMP models, integrate-and-fire models, full HH like models)
% add chapter on information theory, mutual information, stimulus reconstruction, coherence % add chapter on information theory, mutual information, stimulus reconstruction, coherence
% move STA here!
% add a chapter on Bayesian inference (the Neuroscience of it and a % add a chapter on Bayesian inference (the Neuroscience of it and a
% bit of application for statistical problems). % bit of application for statistical problems).